Smith P - Macrophyllum macrophyllum - FAUNAParaguay Handbook of the Mammals of Paraguay Number 27 2008 2005). However the species presence in Misiones Province (Departamentos Capital, San Ignacio and Gral. San Martín), Argentina (Chebez 1996, Massoia et al 2006) very close to the Rio Paraná suggests that it may be more widespread than currently thought. HABITAT: Considered a species of humid forest interior and clearings by Emmons (1999) always in the vicinity of water. López-González (2005) noted that Paraguayan specimens were collected from a small cave over the Arroyo Tacuará, PN Cerro Corá and from the Río Ypané close to Belén, Departamento Concepción where the predominant vegetation type is tropical semi-deciduous forest. ALIMENTATION: Considered the only obligate insectivore amongst the Phyllostomidae (Gardner 2007), this little bat uses its greatly enlarged feet and uropatagium to scoop insect prey from on or near the surface of water (Weinbeer & Meyer 2006). Foraging Behaviour and Diet Following their radiotelemetry study on Barro Colorado Island, Panama Meyer et al (2005) concluded that the species forages exclusively over water - unique amongst the Phyllostomids. Foraging areas ranged from 2.7 to 96.1ha (median 12.3ha for both sexes) and there was high night to night consistency in their use. Core use areas corresponded to approximately 35% of the foraging area and were greater in females (median 5.3ha; range 1.1-54.1ha) than males (median 3.3ha; range 2.6-8.7ha). Davis et al (1964) correctly surmised that the oversized feet were used to take aquatic prey and mentioned insects as a likely prey, but their hypothesis that they may use the feet for "flipping small minnows into the air" has not been borne out by stomach contents data obtained since. The species forages in small groups of 3 to 10 individuals (Weinbeer & Meyer 2006). Wied (1826) found only insect remains in stomachs whilst Harrison & Pendleton (1975) found stomachs from El Salvador to contain nothing other than "finely chewed insect remains". Analysis of the contents of the latter detected the presence of Lepidoptera, Diptera and Coleoptera and it was concluded by Harrison (1975) that flying insects make up the bulk of the prey. Gardner (1977) found water-striders (Hemiptera, Gerridae) in Panamanian stomachs, confirming that prey may be taken from the surface of water. Whitaker & Findley (1980) examined five pellets of the species from Costa Rica and found in order of abundance midges (Chironimidae) 43.3% by volume, Araneida 28.3%, unidentified Coleoptera 15%, unidentified Lepidoptera 4.2%, unidentified Diptera 4.2%, unidentified Hemiptera 3.3% and water-striders (Gerridae) 1.7%. Baud (1989) found microlepidoptera and Trichoptera in the stomach of a male collected in Paraguay on 12 October 1985. Fornes et al (1969) found chitinous remains on the inner side of the uropatagium of Argentine specimens. Quelch (1892) speculated incorrectly that blood-feeding may supplement the diet. Diet in Captivity Under captive conditions Weinbeer & Kalko (2007) observed the foraging behaviour of the species. They demonstrated that prey can be found using only echolocation with a 100% success rate, and proved the importance of echolocation in prey capture by showing experimentally a 0% success rate in capture of prey items located just below the water surface where echolocation could not be used for detection. Bats actively searching for food (the search phase) circled low over the water at a flight speed of 2.95ms -1 (+/-0.8 ms -1 ). Once prey was detected (the approach phase) they switched to a more direct flight style, directing the head, ears and nose leaf towards the prey and reducing speed slightly to 2.48ms -1 (+/-0.6 ms -1 ). Mean detection distance was 1.39m +/-0.3 (range 0.91-1.91m). The terminal phase (including capture of prey) begins with a further reduction in flight speed to 2.22ms -1 (+/-0.4) at a distance of 0.47m +/-0.1 (range 0.32-0.61m). To capture prey the uropatagium is lowered perpendicular to the body stabilized by the long calcars and the head is bent downwards. At this point the claws are maintained parallel to the flight direction either side of the uropatagium and are not lowered into the water (as in Noctilionidae). Upon arriving at the target the distal part of the tail membrane is slid over the surface of the water with only the tips of the calcars touching the water surface and once prey is contacted with the tail membrane the feet are moved together. As a result prey is captured primarily by the tail membrane with the assistance of the claws, and in most cases the claws are not used for removing prey from the water surface. The dermal denticles on the ventral part of the uropatagium probably increase friction for seizing prey, whilst their alignment in longitudinal rows helps reduce water resistance. Following capture the head, ears and nose leaf are returned to the normal position, the head was bent down to remove the prey to the mouth and the bat flew to a perch to consume its meal. During this experiment bats were fed mealworms and each bat consumed 80-100 per night. Smith P 2008 - LONG-LEGGEDBATMacrophyllum macrophyllum - Mammals of Paraguay Nº 27 Page 4

Smith P - Macrophyllum macrophyllum - FAUNAParaguay Handbook of the Mammals of Paraguay Number 27 2008 Bloedel (1955) noted that his captive individual died after only two days without touching the fruit that was offered as food. REPRODUCTIVE BIOLOGY: Seasonality Baud (1989) that a male collected in Departamento Concepción had enlarged testicles (diameter 3.7mm) but that they had not descended to the scrotum. Wilson & Fox (1991) noted that females collected on 10 February 1982 showed signs of recent lactation and some subadults with incomplete fusion of long bones were also present. Rest of range Pregnant females were taken in Guatemala in both the wet and dry seasons, in El Salvador in October, in Costa Rica in March and May and in French Guiana in October and November. (Nowak 1991). Males taken in December in El Salvador were considered sexually active by Felton (1956). Courtship Wilson & Fox (1991) collected one male and 16 females from a small cave on the Arroyo Tacuara in PN Cerro Corrá, Departamento Amambay and suggested that a harem breeding system may operate. Furthermore they cite Felten (1956) who found five males together in a small roost in El Salvador as possible evidence that males without a harem roost together and an unpublished series of specimens collected by Dr CO Handley in Panama and housed at the National Museum of Natural History in Washington which also figured single males with groups of females. An absence of subadult males in the roost that they sampled suggested that young males either disperse or are chased from the harem at an early age. Weinbeer & Meyer (2005) noted that reproductively active males spend less time foraging and more time at the roost than non-reproductive males. Pregnancy Harrison (1975) notes that a foetus measuring 17.5mm taken from a pregnant female in El Salvador during October already possessed feet and claws 8mm long, at that stage of development equal to the forearm length and almost half the body length, and that the dermal denticles were already discernible on the uropatagium. GENERAL BEHAVIOUR: Activity Levels Weinbeer & Meyer (2006) noted maximum activity of the species on Barro Colorado Island, Panama at dusk, when the bats spent about 30 continuous minutes on the wing and that activity continued at a lower, but relatively constant level throughout the night. There was no sexual difference in the amount of time spent in continuous flight (mean 7mins; maximum 60mins), but males spent slightly less time on the wing per night than females (median for males 193.4 v females 207mins). Periods of activity were interrupted by hanging phases at night roosts or in the day roost (median 5.8mins, maximum 3 hours). Bats were found to spend more time in the hanging phase shortly after midnight than they did either at dawn or dusk. Rain reduced or ceased time spent in the air, and though they would continue foraging during drizzle, heavy rain signalled a return to the roost. Emergence time and return to roost were correlated with time of sunset (mean 20.6min +/-1.7min after sunset) and sunrise (mean 30.1min +/-2.5min before sunrise) respectively. Bats tuned their foraging activity to the abundance of aerial prey. Flight Pattern Flight is fast and fluttery, almost butterfly-like. The species is difficult to capture when foraging and most mist-netting captures are at the roost site (Seymour & Dickerman 1982). The species is most frequently captured low over water (Whitaker & Findley 1981) and Davis et al (1964) caught their specimens "in the bottom pocket of the net" within "a foot of the water". Roosts They roost alone or in small groups in caves (López-González 2005), tunnels (Dickerman et al 1981), abandoned buildings (Hill & Bown 1963), ravines (“quebradas” – Ascorra et al 1991), subterranean cavities (Brosset & Charles-Dominique 1990) or under bridges (Gardner 2007). In Panama Bloedel (1955) found colonies to be small and consist of a few individuals, frequently accompanied by Desmodus rotundus. In El Salvador a colony in a culvert was shared with Glossophaga soricina (Harrison & Pendleton 1975). Peracchi & Albuquerque (1971) noticed a roosting association with Glossophaga soricina and Carollia perspicillata in Brazil. Baud (1989) collected a male from a roost at Rio Ypané, Departamento Concepción which was also occupied by Desmodus rotundus, Noctilio albiventris and Platyrrhinus lineatus. Fornes et al (1969) reports that two Argentine specimens were collected in a cave a few metres from the Rio Paraná which had two entrances, a narrow one occupied by a colony of 150 Desmodus rotundus and a wider one where this species was taken with a single Carollia perspicillata. One colony in a 10m long culvert in Brazil contained 50 individuals (Peracchi et al 1984) whilst a colony in culverts at La Avellana, a Guatemalan coastal marsh numbered up to 70 (Dickerman et al 1981). Bats at the Guatemalan roost only used culverts containing water and were absent from dry culverts. They Smith P 2008 - LONG-LEGGEDBATMacrophyllum macrophyllum - Mammals of Paraguay Nº 27 Page 5